Polibotánica

ISSN: 1405-2768
polibotanica@gmail.com
Departamento de Botánica
México

Montejo-Mayo, Wilber; del-Val, Ek; Gomez-Romero, Mariela; de la Barrera, Erick; Lindig-

Cisneros, Roberto
INTERACTIONS BETWEEN DOMINANT HYDROPHYTIC SPECIES OF THE
WETLANDS OF WESTERN MEXICO MEDIATED BY FIRE AND NITRATE
CONCENTRATION
Polibotánica, núm. 40, agosto, 2015, pp. 153-161
Departamento de Botánica
Distrito Federal, México

Available in: http://www.redalyc.org/articulo.oa?id=62142251010

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 Núm. 40, pp. 153-161, ISSN electrónico: 2395-9525; México, 2015
DOI: 10.18387/polibotanica.40.10
INTERACTIONS BETWEEN DOMINANT HYDROPHYTIC
SPECIES OF THE WETLANDS OF WESTERN MEXICO
MEDIATED BY FIRE AND NITRATE CONCENTRATION

INTERACCIONES ENTRE ESPECIES HIDRÓFITAS

DOMINANTES DE HUMEDALES DEL OCCIDENTE DE
MÉXICO MEDIADAS POR EL FUEGO Y LA CONCENTRACIÓN
DE NITRATOS

Wilber Montejo-Mayo1, Ek del-Val2, Mariela Gomez-Romero2,

Erick de la Barrera2, y Roberto Lindig-Cisneros2,3
1
Facultad de Biología, Universidad Michoacana de San Nicolás de Hidalgo, Morelia,
Michoacán, México.
2
Centro de Investigaciones en Ecosistemas, Universidad Nacional Autónoma de México.
Apartado Postal 27, Admón. 3, Santa María, CP 58091, Morelia, Michoacán, México.

SUMMARY complex and differed between sampling

Wetland plant community dynamics are
strongly driven by abiotic factors such as
nutrient levels and fire. In wetlands where
invasive species are present assessing the
role of abiotic factors in plant-plant interac-
tions is fundamental for both understanding
community dynamics and management. In
this study the interaction between pairs of
species was quantified between one non-in-
vasive (Schoenoplectus americanus), one
that can become overdominant after human
disturbances (Typha domingensis) and one
that has an invasive lineage present in North
America (Phragmites australis), growing
under different nitrate concentrations and
subjected to fire. All species responded by
increasing growth to nitrate addition, but
depending on the species, the response was
significant for different variables. Phragmites
for height and root biomass, Typha for aerial
and root biomass and Schoenoplectus for all
variables. Interactions between species were
years. For the first year, only the effect of
Typha on itself was significant. For the se-
cond year, the effect of Typha was negative
on Phrangmites, the effect of Schoenoplectus
was negative on itself and, the effect of
Typha and Phragmites was negative on
Typha. The effect of fire was significant for
Phragmites and Schoenoplectus, not bur-
ned plant were taller. Interactions between
plants were altered by fire, the presence of
Schoenoplectus benefited the performance
of Typha, and the opposite was also true.
For early stages of these species growth,
the strongest competition occurs between
the native species, allowing the invasive
to mostly respond to nitrate levels, and fire
have a strong effect also on the interactions
between the natives and not with the inva-
sive.
Key words: wetland, negative interactions,
management, nutrients, ecological resto-
ration.

153
 Núm. 40: 153-161
RESUMEN
La dinámica de comunidades vegetales de
humedales está influenciada por factores
abióticos, como los nutrimentos y el fue-
go. En humedales en donde hay especies
invasoras, determinar el papel de factores
abióticos en las interacciones planta-planta
es fundamental para entender la dinámica
de la comunidad y para su manejo. En
este estudio, la interacción entre pares
de especies fue cuantificada entre una no
invasora (Schoenoplectus americanus),
una que puede volverse sobredominante
como consecuancia de disturbios huma-
nos (Typha domingensis) y una que posee
un linaje que es invasor en norteamérica
(Phragmites australis), las cuales crecieron
bajo diferentes concentraciones de nitrato y
fueron sometidas al efecto del fuego. Todas
las especies respondieron a la adición de
nitratos incrementando su crecimiento.
Phragmites respondió en altura y biomasa
de raíces, Typha en biomasa aérea y de raíces
y Schoenoplectus para todas las variables.
Las interacciones entre especies fueron
complejas y variaron entre años. Durante
el primer año, sólo el efecto de Typha en
si misma fue significativo. Para el segundo
año, el efecto de Typha fue negativo en
Phrangmites, el efecto de Schoenoplectus
fue negativo en sí mismo y el efecto de
Typha y Phragmites fue negativo en Typha.
El fuego fue significativo para Phragmites
y Schoenoplectus, las plantas no quemadas
fueron más altas. Se detectaron efectos del
fuego en las interacciones, la presencia de
Schoenoplectus benefició el desempeño de
Typha, y viceversa. En etapas tempranas, las
interacciones más fuertes ocurrieron entre
las especies no invasoras, permitiendo a la
invasora responder a la concentración de
nitratos, y el fuego tuvo su mayor efecto
154
Agosto 2015
en la interacción entre nativas y no con la
invasora.
Palabras clave: humedal, interacciones ne-
gativas, manejo, nutrimentos, restauración
ecológica
INTRODUCTION AND METHODS
Wetland plant communities are complex
systems in which abiotic factors can sig-
nificantly alter its floristic composition
(Zedler and Kercher, 2004). Among the most
important of these factors are hydroperiod,
salinity, and nutrient concentration, especially
that of nitrogen and phosphorus (Cronk and
Fennessy, 2001). It has been suggested that
when wetlands are subjected to changes in
the chemical composition of their water
supply, particularly in the form of increases
in nitrogen concentration, changes can occur
in the richness and/or composition of species
that can facilitate the domination of the sys-
tem by invasive species (Cronk, 1995), whe-
ther native or exotic (Zedler and Kercher,
2004). Among the native species that present
invasive behavior in wetlands many belong
to the genus Typha which, in response to
increases in nutrient concentration and other
human disturbances, can form dense monos-
pecific patches and thus displace other native
species (Galatowitsch et al., 1999; Smith et
al., 2001; Woo and Zedler, 2002; Levin et
al., 2006). Others, such as certain lineages
of Phragmites australis (Swearingen and
Saltonall, 2010) are particularly aggressive
invasives (Rickey and Anderson, 2004; Lae-
gue et al., 2006; Saltonstall and Stevenson,
2007). Phragmites australis is a species
considered to be native to Eurasia by some
authors without recognizing subspecies (Lot
et al., 2013), others (Saltonal et al., 2004)
recognize one subspecies that they consi-
Montejo-Mayo, W. et al.: Interactions between dominant hydrophytic species of the Wetlands of Western Mexico
der to be native to Canada and the United
States of America, Phragmites australis
subsp. americanus, and a lineage present
from the Southern United States to Central
America with uncertain origin, although this
last lineage has been proposed as a new
species, Phragmites karka (Ward, 2010).
Despite the taxonomical discrepancies, an
invasive form is recognized, characterized
by morphological and population dynamics
characteristics, in particular that forms dense
clumps that exclude all other plant species
(Sweringen and Saltonall, 2010).
To understand the dynamics of the displace-
ment of native species by invasive species
in wetlands, it is necessary to quantify the
effects of factors, such as nutrient contents
or fire, that can modify the interactions be-
tween species. The present study is based
on observations of the vegetation structure
and dynamics in a model system, the spring
wetlands of the Mintzita (101°17’47” W,
19°38’43” N: 1930 masl) in the municipality
of Morelia, in Michoacán, México (Escutia-
Lara et al., 2009a). These wetlands are cha-
racterized by a natural zonation consisting
of an area dominated by Schoenoplectus
americanus close to the edge of the wetland,
and a zone dominated by Typha domingensis
that conforms to changes in the water level.
Between both of these areas, there is a zone
in which the two species coexist and where
the distributions respond partially to the wa-
ter level throughout the wetland; however,
other factors, such as nutrient concentration
and fire also seem to play an important role.
Phragmites australis forms dense clumps
consistent with those reported for the inva-
sive lineage (Sweringer and Saltonal, 2010)
in the wetlands irrespective of the dominant
native species. In a previous study, it was
found that individuals of T. domingensis in
the wetland respond mainly to the addition
of nitrogen; S. americanus also responds to
increased nitrogen, but also to phosphorus in
terms of the accumulation of aerial biomass
(Escutia-Lara et al., 2010). The responses
of P. australis have not been evaluated in
this system but, in other wetlands, it has
been reported that this species responds
to increases in the concentrations of both
nitrogen and phosphorus and also to the
interaction of these two nutrients (Boar,
1996; Ennabili et al., 1998; Ehrenfeld,
2003). From field observations in the model
wetland (Escutia-Lara et al., 2009a) and
from controlled experiments (Escutia-Lara
et al., 2009b; Escutia-Lara et al., 2010), it
can be suggested that the competitive capa-
city of P. australis is greater or equal to that
of T. domingensis and, in turn, greater than
that of S. americanus; however, the mag-
nitude and exact nature of the interactions
between these species remains unknown.
The objectives of the present study were
to quantify the response of these species to
different concentrations of nitrate, as this is
the macronutrient to which all three species
respond most markedly (Escutia-Lara et
al., 2010) and to fire, in order to relate this
recurring disturbance factor to patterns of
species growth.
Rhizomes of the three species were collected
at the study site; these were selected to be of
similar size at the start of the experiment and
were assigned at random to each of the re-
plicates of the experiments described below.
Experiments were conducted to quantify
the effect of interactions between pairs of
species: Typha domingensis, Phragmites
australis and Schoenoplectus americanus.
Each experimental unit consisted of two
plants, either of the same species or of each
possible pairing between them. Rhizomes
155
 Núm. 40: 153-161
were planted in five liter pots (25 cm high
x 16 cm wide) filled with commercial peat.
Thirty pots were planted with each one of
the six possible species combinations (three
with interspecific and three with intraspeci-
fic pairs) giving a total of 180 experimental
units. In addition, for each species-interac-
tion treatment, the effect of five treatments
of fertilization with potassium nitrate
(KNO3) was tested: 0, 0.08, 0.16, 0.24 and
0.32 g/l of cultivation medium. In this way,
six replicates were produced for each treat-
ment of the combination of competition x
fertilization. Addition of KNO3 was carried
out every 15 days for a period of six months
(May to October). During the dry season
(February of the following year), 15 pots
were chosen at random from each interac-
tion treatment (three for each fertilization
X interaction combination of treatments)
in order to burn the aerial biomass, while
the other 15 were left as a control. In June,
all remaining plants were harvested and
aerial and root biomass recorded. In order
to assess the importance of the interaction
between the different species, three response
variables were analyzed: plant height, and
aerial and root biomass. In the case of same
species interactions, the response variable
considered was the average value from
both plants in each pot. For different species
interactions, the variable was measured in
the individual of the species on which the
effect of the interaction was to be determi-
ned. Analysis was conducted with general
linear models, considering fertilization as a
continuous variable, and the interaction and
aerial biomass burning as categorical varia-
bles. Tests of hypothesis were carried out to
generate ANOVA tables (Crawley, 2007)
and to determine the effect of each of the
variables and their interactions. All analyses
156
Agosto 2015
were conducted with the statistical program
R (R Development Core Team, 2011).
RESULTS
Following burning of the aerial parts of the
plants in the second growing season, all
species responded to the addition of nitrate
and, in some cases, to the interaction treat-
ments and burning. For P. australis, there
was a significant response in plant height
to the added fertilizer (fig. 1a; F(1,78) =
71.2, P < 0.000001). The effect of the inte-
ractions (fig. 1b) was significant (F(2,78) =
6.6, P = 0.002): the intraspecific interaction
produced the greatest average plant height
(138 cm), followed by the interaction with
S. americanus (120 cm) and with T. do-
mingensis (115 cm). Burning of the aerial
parts did not affect height in P. australis.
In the case of aerial biomass (fig. 1b), sig-
nificant differences were observed due to
the addition of nitrate (F(1,78) = 118.3, P <
0.000001) and to the effect of the interaction
(F(2,78) = 6.9, P = 0.002). These followed
the same pattern observed in plant height
(41 g for intraspecific interaction, 32 g for
the interaction with S. americanus and 26
g for the interaction with T. domingensis).
Significant differences (F(1,78) = 8.32,
P = 0.005) were found with the burning
treatment, since greater quantities of aerial
biomass accumulated in those plants that
had not been burned (36 g), relative to those
that had (30 g). Root biomass (fig. 1c) only
increased with increased nitrate addition
(F(1,78) = 18.4, P < 0.0001).
Schoenoplectus americanus presented a
more complex set of responses than P.
australis. In terms of height (fig. 1d), this
species responded to both nitrate addition
Montejo-Mayo, W. et al.: Interactions between dominant hydrophytic species of the Wetlands of Western Mexico

Fig. 1. Response of Phragmites australis, Schoenoplectus americanus and Typha domingensis

to the treatments of nitrate addition, competition and burning at the end of the experiment.
Each panel (a to h) shows the relationship between the response variable and the con-
centration of added nitrate, and the means and standard errors for the response variable
as a function of the competition interaction or the burning treatments. Slope curves are
shown only when the effect of the nitrate addition was significant. In cases where some of
the interactions between factors were significant, adjustment curves are shown for each
combination of levels. Different letters denote significant differences in means according
to the Tukey test.

157
 Núm. 40: 153-161
(F(1,78) = 99.1, P < 0.000001) and to the
interaction with other individuals (F(2,78)
= 3.24, P = 0.045), since the plants reached
the tallest average height (95 cm) in the
presence of P. australis, followed by those
grown with T. domingensis (92 cm) and
finally by those grown in the intraspecific
treatment (86 cm). They also responded
to the burning treatment (F(1,78) = 9.1, P
= 0.003), with plants that had been burned
reaching an average height of 96 cm, while
non-burned plants only reached 86 cm.
There were also significant effects of the
interaction between nitrate addition and bur-
ning treatment (F(1,78) = 4.15, P = 0.045),
since non-burned plants presented lower
rates of height increase than burned plants,
with increasing concentrations of nitrates, as
reflected in the slopes of the curves. The in-
teraction between competition and burning
treatment was also significant (F(2,78) =
5.73, P < 0.004) because the burned plants
from the treatment of competition with T.
domingensis reached a greater height (104
cm) than any other combination of levels
of the two factors while non-burned plants
of the same competition treatment were the
shortest (80 cm).
Regarding aerial biomass (fig. 1e), S.
americanus responded to the addition of
nitrate (F(1,78) = 43.1, P < 0.000001) and
to competition (F(2,78) = 56.2, P < 0.001),
accumulating more biomass in the inters-
pecific treatment with P. australis (4.0g)
relative to the other two treatments (2.4 g
with T. domingensis and 2.3 g in intraspe-
cific competition). In the burning treatment,
more aerial biomass was accumulated in the
non-burned plants (3.35 g) than in burned
plants (2.5 g). Regarding root biomass (fig.
2f), S. americanus presented the most com-
158
Agosto 2015
plex pattern of responses to treatments: there
were significant effects of nitrate addition
(F(1,78) = 91.0, P < 0.000001), plant-plant
interactions (F(2,78) = 8.0, P = 0.0006) and
of the combination of factors (F(2,78) = 4.1,
P = 0.02), since the intraspecific interaction
treatment with fire produced a lower root
biomass (4.6 g). Finally, the interaction be-
tween the three factors; addition of nitrate,
plant-plant interactions and burning was
also significant (F(2,78) = 5.5, P = 0.005),
as reflected in the slopes of the curves of
root biomass as a function of nitrate addi-
tion for each combination of the levels of
experimental factors.
Typha domingensis responded to fewer
factors than the other two species, since
for no variable was there a response to the
burning treatment. For plant height (fig. 1g),
this species responded to the addition of
nitrate (F(1,78) = 17.0, P < 0.0001) and to
plant-plant interactions (F(2,78) = 18.3, P <
0.000001) since the plants reached greatest
height on interaction with S. americanus
(137 cm) and presented the shortest height
with P. australis (118 cm). Accumulation
of aerial biomass (fig. 1h) also responded to
nitrate addition (F(1,78) = 12.0, P < 0.0008),
and to plant-plant interactions (F(2,78) =
14.5, P < 0.0001), since the plants in intras-
pecific treatments accumulated the least
biomass on average (3 g), followed by plants
grown with P. australis (5.8 g) and finally
those grown with S. americanus (9.0 g);
the accumulation of root biomass only res-
ponded to plant-plant interactions (F(2,78)
= 7.5, P = 0.001), and followed the same
pattern observed in the aerial biomass; 11.8
g in intraspecific competition, 14.1 g with
P. australis and 23.9 g with S. americanus).
Montejo-Mayo, W. et al.: Interactions between dominant hydrophytic species of the Wetlands of Western Mexico
DISCUSSION
The results show, as expected, that all the
study species responded positively to the
addition of nitrate; however, in the pre-
sence of other individuals, performance
variables differed depending on species.
Interspecific interactions are not important
for Phragmites australis and Schoenoplectus
americanus since they present the same
rates of growth as when they are growing
alongside individuals of their own species.
For Typha domingensis, the situation is
different since intraspecific interactions
were more detrimental than interspecific
ones. Given that P. australis is invading the
wetland, it was expected that P. australis
would be a superior competitor to both T.
domingensis and S. americanus and would
suppress their performance, but this did not
in fact occur. The addition of nutrients alone
cannot explain the increase in abundance
of P. australis in the Mintzita wetland, as
nutrients do for other invasive species in
other wetlands (Woo and Zedler, 2002). In
the case of other species that present inva-
sive behavior, it has been documented that
they require anthropogenic disturbances in
order to increase their numbers (Jesson et
al., 2000; Fireswyck and Zedler, 2007).
In this experiment, the added disturbance
is related to the burning of vegetation, a
practice that currently takes place in Mint-
zita (Escutia-Lara et al., 2009a). With this
disturbance, the plant response to nutrients
remained surprisingly unchanged; the three
species grew in line with the increases in
nitrate concentration. However, the inter-
relations associated with plant-plant inter-
actions did change. For burned P. australis
plants, interspecific interactions caused
diminished growth, since taller and more
robust plants were produced when grown
in the presence of individuals of the same
species. For T. domingensis, the contrary
was true as the presence of S. americanus
benefited its performance after burning and
for S. americanus, the plants most favored
were those grown with T. domingensis. The
positive effects of ash from S. americanus in
the growth of T. domingensis seedlings have
been reported (Lopez-Arcos et al., 2012). It
therefore appears that competition between
these two species is less when growing to-
gether and there is a likelihood of fire. The
modification of these competitive interrela-
tions by anthropogenic disturbance, reported
in previous studies for other wetland species
(Woo and Zedler, 2002) as being of benefit
to those species with invasive attributes, was
not reflected in this experiment. It is worth
noting that P. australis is always taller and of
greater biomass than the other two species,
making it likely in the long term that compe-
tition for light becomes a determining factor
in its invasion of the site, and therefore this is
a factor that must be fully taken into account
in any future studies to predict the behavior
of this invasive species.
CONCLUSIONS
The series of experiments indicate that nitrate
availability altered the intensity of plant-
plant interactions but that fire was more
instrumental in altering these, in particular
that presence of Schoenoplectus benefited
Typha. Since several species of the later
genus have shown invasive behavior, this
study suggests an additional mechanism that
deserves more attention under natural condi-
tions, since the controlled conditions of the
experiments undertaken limits its realism.
159
 Núm. 40: 153-161
Finally, fire was a significant factor altering
growth for Phragmites and Schoenoplectus,
benefiting more the former than the later,
therefore, strongest competition occurred
between natives, leaving Phragmites, the
invasive, free to respond to increased nitrate
availability by growing more.
A CKNOWLEDGEMENTS
This work was funded by the Universidad
Nacional Autónoma de México through the
project PAPIIT IN203608 and CONACYT
(SEP-CONACYT-2008-101335).
LITERATURE CITED
Boar, R.R., 1996. “Temporal variations in
the nitrogen content of Phragmites
australis (Cav) Trin ex Steud from
a shallow fertile lake”. Aq. Bot., 55:
171-181.
Crawley, R., 2007. The R book. John Wiley
and Sons, Ltd., USA.
Cronk, J. K., y M.S. Fennessy, 2001. Wet-
land Plants: Biology and Ecology.
Lewis Publishers. Washington D.C.
EUA. 462 pp.
Cronk, Q.C.B., 1995. Plant Invaders. Cha-
pman and Hall. London, UK.
Ehrenfeld, J.G., 2003. “Effects of exotic
plant invasions on soil nutrient cycling
processes”. Ecosystems, 6: 503-523.
Ennabili, A.; M. Ater, y M. Radoux, 1998.
“Biomass production and NPK reten-
tion in macrophytes from wetlands
of the Tingitan Peninsula”. Aq. Bot.,
62: 45-56.
160
Agosto 2015
Escutia-Lara, Y.; E. de la Barrera, Y. Martí-
nez de la Cruz, y R. Lindig-Cisneros,
2010. “Respuesta a la adición de ni-
trógeno y fósforo en el crecimiento de
Typha domingensis y Schoenoplectus
americanus”. Boletín de la Sociedad
Botánica de México, 87: 93-97.
Escutia-Lara, Y.; M. Gómez-Romero, y R.
Lindig-Cisneros, 2009b. “Nitrogen
and phosphorus effect on Typha do-
mingensis Presl. rhizome growth in a
matrix of Schoenoplectus americanus
(Pers.) Volkart ex Schinz and Keller”.
Aq. Bot., 90: 74-77
Escutia-Lara, Y.; S. Lara-Cabrera, y R.
Lindig-Cisneros, 2009a. “Fuego y
dinámica de las hidrófitas emergentes
del humedal de la Mintzita, Michoa-
cán, México”. Revista Mexicana de
Biodiversidad, 80: 771-778.
Frieswyk, C.B., y J.B. Zedler, 1997. Vege-
tation Change in Great Lakes Coastal
Wetlands: Deviation from the Histo-
rical Cycle. J. Great Lakes Res., 33:
366-38.
Galatowitsch, S.M.; N.O. Anderson, y P.D.
Ascher. 1999. “Invasiveness in wet-
land plants in temperate North Ame-
rica”. Wetlands, 19: 733-755.
Jesson, L.; D. Kelly, y A. Sparrrow, 2000.
The importance of dispersal, disturban-
ce, and competition for exotic plant
invasions in Arthur’s Pass National
Park, New Zealand. New Zealand J.
Bot., 38: 451-468.
League, M.T.; E.P. Colbert, y D.M. Seliskar,
2006. “Rhizome growth dynamics
Montejo-Mayo, W. et al.: Interactions between dominant hydrophytic species of the Wetlands of Western Mexico
of native and exotic haplotypes of
Phragmites australis (common reed)”.
Estuaries and Coasts, 29: 269-276.
Levin, L.A.; C. Neira, y E.D. Grosholz,
2006. “Invasive cordgrass modifies
wetland trophic function”. Ecology,
87: 419-432.
Lopez-Arcos, D.; M. Gomez-Romero, R.
Lindig-Cisneros, y P.H. Zedler, 2012.
“Fire-mobilized nutrients from hydro-
phyte leaves favor differentially Typha
domingensis seedling growth”. Env.
Exp. Bot., 78: 33-38.
Lot, A.; R. Medina Lemos, y F. Chiang,
2013. Plantas acuáticas mexicanas:
Una contribución a la flora de México.
vol. I, Monocotiledóneas. Instituto de
Biología. Universidad Nacional Autó-
noma de México.
R Development Core Team, 2011. “R:
A Language and Environment for
Statistical Computing. R Foundation
for Statistical Computing”. Vienna,
Austria. ISBN 3-900051-07-0. URL:
http://www.R-project.org.
Rickey, M.A., y R.C. Anderson, 2004.
“Effects of nitrogen addition on the
invasive grass Phragmites australis
and a native competitor Spartina pec-
tinata”. J. Appl. Ecol., 41: 888-896.
Saltonall, K.; P.M. Peterson, y R.J. Soreng,
2004. “Recognition of Phragmites
australis subsp. americanus (Poa-
Recibido: 17 octubre 2013. Aceptado: 23 enero 2015.
ceae:Arundinoideae) in North Ameri-
ca: Evidence from morphological and
genetic analyses”. Sida, 21: 683-692.
Saltonstall, K., y J.C. Stevenson, 2007. “The
effect of nutrients on seedling growth
of native and introduced Phragmites
australis”. Aq. Bot., 86: 331-336.
Smith, S.M.; S. Newman, P.B. Garret, y
J.A. Leeds, 2001. “Differential effects
of surface and peat fires on soil cons-
tituents in a degraded wetland of the
Northern Florida Everglades”. J. En-
viron. Qual., 30: 1998-2005.
Swearingen, J., y K. Saltonstall, 2010.
“Phragmites Field Guide: Distinguishing
Native and Exotic Forms of Common
Reed (Phragmites australis) in the
United States”. Plant Conservation
Alliance, Weeds Gone Wild. http://
www.nps.gov/plants/alien/pubs/index.
htm.
Ward, D.B., 2010. “North America Has Two
Species of Phragmites (Gramineae)”.
Castanea, 75: 394-401.
Woo, I., y J.B. Zedler, 2002. “Can nutrients
alone shift a sedge meadow towards
dominance by the invasive Thypa X
glauca?”. Wetlands, 22: 509-521.
Zedler, J.B., y S. Kercher, 2004. “Causes
and Consequences of Invasive Plants
in Wetlands: Opportunities, Opportu-
nists, and Outcomes”. Critical Revies
in Plant Sciences, 23: 431-452.
161